LECTURE XII

INTRA-SELECTION OR SELECTION AMONG TISSUES

Does the Lamarckian principle really play a part in the transformations of species?—Darwin's position in regard to this question—Doubts expressed by Galton and others—Neo-Lamarckians and Neo-Darwinians—Results of exercise and practice: functional adaptation—Wilhelm Roux,Kampf der Theile.

Wehave devoted a whole series of lectures to studying the Darwin-Wallace principle of Natural Selection and the range of its operation. It seemed to us to make innumerable adaptations intelligible up to a certain point. We now understand how the purposefulness, which we meet with everywhere among organisms, can have arisen without the direct interference of a Power working intentionally towards an end—simply as the outcome and result of the survival of the fittest. The two forms of the processes of selection, 'natural selection' in the narrower sense, and 'sexual selection,' dominate, so to speak, all parts and all functions of the organism, and are striving to adapt these as well as possible to the conditions of their life. And although the range of operation of Natural Selection is incomparably greater, because it actually affects every part, yet we must attribute to sexual selection also, at least among animals, a range of influence by no means unimportant, since through it, as far as we can see at present, not only do the secondary sexual characters in all their diversity arise, but by the transference of these to the other sex that too is modified, and thus the whole species may be influenced, and may indeed be started afresh on an unlimited series of further transformations.

But although the processes of selection play such an important part in the transformations of the forms of life, we have to inquire whether they are thesolefactors in these transformations, whether the accumulation of chance variations in the direction of utility has been the sole factor in bringing about the evolution of the animate world, or whether other factors have not also co-operated with it.

We are all aware that Lamarck regarded the direct influence of use and disuse as the most essential factor in transformation, and that Darwin, though hesitatingly and cautiously, recognized and accepted this factor, which he believed to be indispensable. Indeed, it seemsat first sight to be so. There is a whole range of facts which seem to be intelligible only in terms of the Lamarckian theory; in particular, the existence of numberless vestigial or rudimentary organs which have degenerated through disuse, the remains of eyes in animals which live in darkness, of wings in running birds, of hind legs in swimming mammals (whales), and of ear muscles in Man, who no longer points his ears, and so forth through a long list.

According to Wiedersheim, there are in Man alone about two hundred of these vestigial or rudimentary organs, and there is no higher animal which does not possess some. In all, therefore, a piece of the past history of the species is embodied in the actually existing organism, and bears witness to the fact that much of what the ancestors possessed is now superfluous, and is either transformed, or is gradually set aside, or is still in process of being set aside. It seems obvious that this gradual dwindling and degeneration of an organ no longer needed cannot be explained through natural selection in the Darwin-Wallace sense, for the process goes on so exceedingly slowly that the minute differences in the size of an organ, which may occur among individuals of the species at any given time during the retrogressive process, cannot possibly have a selection value. Whether the degenerate and now functionless hind leg of the whale is a little larger or a little smaller can have no importance in the struggle for existence; the smaller organ cannot be considered either as a lesser hindrance in swimming or as a greater economy of material, and the case is the same in regard to most other instances of degeneration through disuse. We therefore require another interpretation, and at first sight this seems to be supplied by the Lamarckian principle.

But the reverse process, the strengthening, the enlarging, and the more perfect development of a part, very often goes on proportionately to its more frequent use, and here again the Lamarckian principle seems to afford a simple explanation. For we know that exercise strengthens a part, as disuse weakens it, and if we could assume that these results of use and disuse were transmitted from the individual who brought them about or 'acquired' them in the course of his life to his offspring, then there would be nothing to object to in the Lamarckian principle. But it is precisely here that the difficulty lies. Can we assume such a transmission of 'acquired' characters? Does it exist? Can it be demonstrated?

That Lamarck did not even put this question to himself, but assumed such transmission as a matter of course, is readily intelligible when we consider the time at which he lived. He was himself one of the first to grasp the idea of the transmutation-hypothesis, and hewas only too glad to have any sort of principle of interpretation ready to work with. But Charles Darwin, too, attributed a not inconsiderable influence to this principle, although the transmission of 'acquired' characters which it took for granted was not accepted without reflective hesitation. He even directed his own particular theory of heredity, as we shall see, especially to the explanation of this supposed form of inheritance, and we can very well understand this, after what I have said as to the impossibility of explaining the disappearance of organs which have become superfluous by the Darwin-Wallace theory of Natural Selection. Darwin needed the Lamarckian principle for the explanation of these phenomena, and it was this that decided him to assume the transmission of 'acquired' characters, although the proofs of it can hardly have satisfied him. For when we are confronted with facts which we see no possibility of understanding save on a single hypothesis, even though it be an undemonstrable one, we are naturally led to accept the hypothesis, at least until a better one can be found. It is in this way, obviously, that we are to understand Darwin's attitude to the Lamarckian principle; he did not reject it, because it seemed to him to offer the only possible explanation of the disappearance of characters which have become useless; he adhered to it, although the transmission of acquired characters which it assumed must have seemed, and, in point of fact, did seem to him doubtful, or at least not definitely proved. Doubts, some faint, some stronger, as to this assumed form of inheritance were hardly expressed till somewhat late in the day—almost twenty years after the appearance of theOrigin of Species—first by Francis Galton (1875), then by His, who definitely declared himself at least against any inheritance of mutilations, and by Du Bois-Reymond, who, in his addressUeber die Uebungin 1881, said: 'If we are to be honest, we must admit that the inheritance of acquired characters is a hypothesis inferred solely from the facts which have to be explained, and that it is in itself quite obscure.'

This is how it must appear to every one who examines it simply in respect of its theoretical possibility, its conceivability. This is how it appeared to me when I attempted, in 1883, to arrive at clearness on the subject, and I then expressed my conviction that such a form of inheritance was not only unproved, but that it was even theoretically unthinkable, and that we ought to try to explain the fact of the disappearance of disused parts in some other way, and I attempted to give an explanation, as will be seen later.

Thus war was declared against the Lamarckian principle of the direct effect of use and disuse, and there arose a strife whichhas continued down to the present day, the strife between the Neo-Lamarckians and the Neo-Darwinians, as the two disputing parties have been called.

In order to form an independent opinion in regard to this famous dispute, it is, first of all, necessary to examine what actually takes place when an organ is exercised or is left inactive, and further, whether we can assume that the results of this exercise or inaction can be transmitted to descendants.

That exercise in general has a strengthening, and neglect of it a weakening influence on the relevant organ has long been known and is familiar to all; gymnastics make the muscles stronger, the thickness of the exercised muscle and the number of its fibres increases; the right arm, which is much more used than the left, is capable of performing twenty per cent. more work. Similarly, the activity of glands is increased by exercise, and the glands themselves are increased in size, as are the milk-glands of the cow through frequent milking; and that even the nerve-elements can be favourably influenced by exercise is proved by actors and professors of mnemonics, who have by practice increased their powers of memory to an almost incredible degree. I have heard of a singer who had learned by heart 160 operas; and which of us has not experienced how quickly the capacity for learning by rote can be again increased by practice, even after it has been neglected or left unexercised for a long time?

I have always been particularly struck with the practising of a piece of music, with its long succession of periods of different phrase, with its changes in melody, rhythm, and harmony, which nevertheless becomes so firmly stamped on the memory that it can be played, not only consciously, but quite unconsciously, when the player is thinking intensely of other things. It is in this case not the memory alone, but the whole complicated mechanism of successive muscle-impulses, with all the details of fast and slow, loud and soft, that is engraved on the brain elements, just like a long series of reflex movements which set one another a-going. Though in this case we cannot demonstrate the material changes which have taken place in the nervous elements, there can be no doubt that changes have taken place, and that these consist in a strengthening of definite elements and parts of elements. The strengthening causes certain ganglion-cells to give a stronger impulse in a particular direction, and this impulse acquires increasing transmissive power, and so on.

Our first theoretical insight into these relations came through Wilhelm Roux, who, in 1881, gave expression to what had previouslybeen an open, if not quite conscious, secret, that 'functional stimulus strengthens the organ,' that is to say, that an organ increases through its own specific activity. Up till that time it had been believed that it was merely the increased flow of blood that caused the increase in the size of a much-used part. Roux showed that there is a 'quantitative self-regulation of the organ according to the strength of the stimulus supplied to it'; that the stimulated organ, that is, the organ which is performing its normal function, may, in spite of the increased breaking down or combustion (dissimilation), assimilate all the more rapidly; that its used-up material is 'over-compensated,' and that therefore it grows. He called this the 'trophic' or nutritive effect of the stimulus, and in terms of this he explained the increase and the heightened functional capacity of the much-used organ. Conversely, he referred the decrease of a disused organ to 'functional atrophy,' which sets in when there is a deficient compensation for the substance used up in the metabolism.

But if we press for deeper analysis, we must ask: 'On what does this trophic effect of functional stimulus depend?' Roux could not answer this question when he wrote, nor can we do so now, as Plate has justly emphasized. We are here face to face with the fundamental phenomenon of life, metabolism; and, since we do not understand the causes of this, we are not in a position to say why it varies in this way or in that according to the 'stimulus.' But the fact itself is certain that the organs respond up to a certain point to the claims made upon them; they increase in proportion as they function more frequently or more vigorously, they are able to respond to increased functional demands, and this Roux has called 'functional adaptation.' As an animal adapts itself to the claims of the conditions of its life, for instance, by taking on a green or a brown protective colour according as it lives on green or brown parts of plants, so the individual organ adapts itself to the strength of the stimulus which impels it to function, and increases or decreases in proportion to it. Ifonekidney in Man degenerate, or be surgically removed, the other begins to grow, and goes on increasing until it has reached nearly twice its former size. The specific stimulus which is brought to bear upon it by the urea contained in the blood, and which forces it to grow, is twice as great in the absence of the other kidney, and therefore the remaining kidney grows in response to the increased stimulus and its 'trophic effect' until its increase in size has reduced the functional intensity to the normal proportion.

Adaptation of an organ in the opposite direction takes place when the function diminishes or ceases. If a nerve supplying amuscle or a gland be cut through, the organ concerned begins to degenerate and to lose its normal structure to a greater or less degree. Sensory nerves also degenerate in their peripheral part when they are cut through. In such cases there may be no alteration either in the nutritive mechanism or in the blood-vessels, &c., but the functional stimulus—in the case of the muscle, the stimulus from the will—no longer affects the organ, and its metabolism is so much lowered in consequence that it begins to degenerate.

When we admit that the fit adaptation of the organism, as far as we understand it, must depend upon processes of selection, we may refer this 'functional adaptation' also to primitive processes of selection, which prevailed at the very beginning of life upon our earth, and represented, so to speak, the first adaptation that was established, but we can say nothing with certainty in regard to this matter as long as we do not understand the essence of assimilation. It is conceivable, however, that aprimaryadaptiveness may have arisen, so to speak, abruptly, through a concurrence of favourable circumstances, as we shall endeavour to show later on when we discuss the beginnings of life.

Even although we cannot lay bare the primary roots of 'functional adaptation' we can gain from the fact itself very valuable insight into phenomena which would otherwise be unintelligible and mysterious:the perfectly adapted structure of many tissues and their power of adaptation to changed conditions. In this lies, in the main, the advance in our knowledge which is due to Roux'sKampf der Theile.

If a number of embryonic cells of different capacity, sayA,B, andC, be affected by different kinds of functional stimuli,a,b, andc, those cells will grow most rapidly which are most frequently affected by the stimulus appropriate to them. The proportion in which the cellsA,B, andCwill ultimately be present in the tissues will depend upon the frequency with which the stimulia,b, andcact upon the tissue. But the tissue will be still more precisely determined as to its structure if the three kinds of stimuli affect the cell-mass, not uniformly all over, but only at certain spots, or along particular paths, one in this, the other in that. Thus the cellsAwill predominate over the cellsBandCat all the places which are most frequently affected by the stimulusa, the cellsBin the sphere of the stimulusb, and the cellsCin that of the stimulusc; there they will increase most rapidly and so crowd out the other kinds of cells, and thus a spatial arrangement will be established within the tissue, a 'structure' which corresponds and is well adapted to its end. This is what Roux deduced from hisStruggle of the Parts, and I subsequently defined the process as histonal or tissue selection.

Let us first take an example. The anatomist Hermann Meyer showed in 1869 that the so-called 'spongiosa,' that is, the bony tissue of spongy structure within the terminal portions of the long bones in Man and Mammals, has a minute structure conspicuously well adapted to its office. The thin bone lamellæ of this 'spongiosa' lie precisely in the direction of the strongest strain or pressure which is exerted upon the bone at the particular area. Arch-like in form, they are kept apart by means of buttresses, and no architect could have done better if he had been entrusted with the task of making a complicated system of arches with the greatest possible carrying and resisting power combined with the greatest possible economy of material.

This well-adapted structure is now interpreted through theStruggle of the Partsas a self-differentiation, for if there be in the rudiments or primordia of the bone differently endowed elements[10], that is, cells which respond in diverse ways to different stimuli, these must arrange themselves locally, owing to the struggle for space and food, in a manner corresponding to the distribution of the different stimuli in the bone. The largest amount of bone substance will be formed in the directions of the strongest strain and the greatest pressure, because the bone-forming cells are excited by this, their functional stimulus, to growth and multiplication. Thus the buttress and arch structure comes about, and between the delicate bone lamellæ spaces will remain free, and these, being relieved from the burden of strain and pressure by the aforesaid bony lamellæ, will offer suitable conditions of life to cells with other functional properties, such as connective tissue cells or vascular cells.

[10]I do not here enter into the question whether we have not in this case to do with similar elements, which have the power of differentiating into one or another kind of cell according to the nature of the external stimuli by which they are influenced.

The structure of the bone spongiosa is not everywhere the same, and it is demonstrably related with precision to the conditions of strain and pressure at each particular region. Thus, just below the soft cartilaginous covering of the joints there are no long pillars with short arches, but only rounded meshes, because the pressure is here almost equally strong from all sides. The long parallel pillars only occur further down in the bone, and they lie in two directions which intersect each other obliquely, corresponding to the two main directions of pressure. But it is only under the functional stimulus of pressure that the bone-forming cells have an advantage over theothers, and multiply more quickly, thus crowding out those that are not attuned to the appropriate functional stimulus.

In a similar manner Roux interprets, in the light of the struggle of the parts, the striking adaptations in the course, the branching, and the lumen-formation of the blood-vessels, in the direction of the intersecting connective tissue strands in the tail-fin of the dolphin, in the direction of the fibres in the tympanum, and in many other adaptations in the histological structure of complex tissues.

In this there is manifestly an important step of progress, for it is obvious that the direction of the bone-lamellæ and such like could not have been determined by individual selection, and the same is true in regard to many other histological details. It cannot be disputed, however, that there is a kind of selection-process here also, similar to that which we think of, with Darwin and Wallace, as occurring between individual organisms. Just as in the latter, which we shall henceforward callpersonal selection, variability and inheritance lead, in the struggle for existence, to the survival of the fittest, so, in histonal differentiation, the same three factors lead to the victory of what is best suited to the parts of the body in question. The tissues and the parts of the tissues have to distribute and arrange themselves so that each comes to fill the place in which it is most effectively and frequently affected by its specific stimulus, that is, the stimulus in regard to which it is superior to other parts; but these places are also those the occupation of which by the best re-acting parts makes the whole tissue capable of more effective function, and therefore makes its structure the fittest. Variability—in this case that of embryonic cells, with different primary constituents—must be assumed; inheritance is implied in the multiplication of the cells by division; and the 'struggle for existence' here assumes its frequent form of a competition for food and space; the cells which assimilate more rapidly because of the more frequent functional stimulus increase more rapidly, draw away nourishment from the more slowly-multiplying cells around them, and thus crowd these out to a greater or less extent.

We might even speak of histonal selection among unicellulars, for it is conceivable that in primitive living substance, such as that of a moneron, there may be minute differences among the vital particles, involving also functional distinctions, which, under the influence of diverse stimuli, may gradually give rise to an increasingly complex differentiation. For the variations in the primary living substance most strongly affected by a particular stimulus would tend to accumulate at the places most frequently reached by that stimulus, andwould crowd out other variations at that spot, just as the body and its individual parts may be said to have taken their architectural form in exact response to the demands made upon them by function. In this case, of course, personal selection and histonal selection co-operate, for every improvement in the organization of the fundamental living substance means at the same time a lasting improvement in the whole individual.

In many-celled organisms, however, we must admit that there is an essential difference between personal and histonal selection, inasmuch as the latter can give rise to adaptive structural modifications corresponding to the needs of the tissue at the moment, but not to permanent and cumulative changes in the individual elements of the tissue. If a broken bone heals crookedly, the spongy substance within the healed portion does not remain as it was before, for the pillars and arches, which now no longer run in the direction best suited to their function, break up, and a new system of arches is formed, not in line with the earlier one, but adapted to the new conditions of pressure. This is certainly an adaptation through selection, but the elements, that is the cells which form the bone substance in response to strain and pressure, or those which in response to the stimulus of the blood flowing into the spaces form the blood-vessels, or those which being quite freed from one-sided pressure develop into connective tissue, must be presupposed. These kinds of cells must be virtually implied in the germ-rudiment; they are themselves adaptations of the organism, and can therefore only be referred topersonal selection. And this is true of all adaptations of theelementsof multicellular organisms, and thus of thecells. Their adaptation according to the principle of division of labour, their differentiation into muscle, nerve, and gland cells can only be referred to natural selection in the Darwin-Wallace sense, and cannot depend upon histonal selection. In the spongy substance of the bone a better bone-cell does not struggle with an inferior one and leave behind it by its survival a host of descendants which are, if possible, better than itself; the struggle for existence and for descendants, in this case, is between two kinds of cell which were different from the beginning, and of which one has the advantage at one spot, another at another. The case may be compared to that of a flock of nearly allied species of bird, of which one species thrives best in the plains, another among the hills, and a third among the mountain forests, all mingled together in a vast new territory to which they had migrated, and in which all three kinds of conditions were represented. A struggle would arise among the different species, in which in everycase the particular species would be victorious which was best adapted to the local conditions. But each would thrive best in the region in which it was superior to the others, and very soon the three species would be distributed as they were in the land from which they came—in the plains, the high lands, and the mountain forests. This would be the result of a struggle between the three species,not between individuals within each species, and it could not therefore bring about an improvement of a single species, but only the local prevalence of one or another. The characters which made one species adapted for the plain, another for the mountain forest werealready there; they can only be referred to personal selection, which brought about the adaptation of their ancestors in the course of ages to the conditions of their life. Something similar is true of the adaptations of the tissues; the differentiation of the individual kinds of cells is an ancient inheritance, and depends upon personal selection, but their distribution and arrangement into specially adapted tissues, so far as there is any plasticity at all, depends upon histonal selection. Obviously, however, only as far as the tissue is plastic, that is, with the power of adjusting itself to particular local conditions. Only adaptations of this kind can be referred to histonal selection; the ground-plan, even of the most complicated tissue, such as the large glands of mammals, the kidneys, the liver, and so on, must have been implicit in the germ, and must therefore be referred to personal selection. A precise limitation of the respective spheres of action of personal selection and histonal selection is not possible as yet, since hardly any investigations on the subject are available.

Roux undoubtedly over-estimated the influence of his 'struggle of parts' when he believed that the most delicate adaptations of the different kinds of cells depended on it. I admit that, for a considerable time, I made the same mistake, until it became clear to me, as it did first in regard to the sex-cells, that this is not, and cannot be the case. How, for instance, could the diverse and minutely detailed adaptations of the sex-cells—which we are to discuss in a subsequent lecture—have arisen in this way? As far as the individual sperm-cell is concerned, it is a matter of indifference whether its head is a little thinner or thicker, its point a little sharper or blunter, its tail a little stronger or weaker. This does not decide whether the cell is to thrive better, or to occur in greater numbers than some other variety. But it does decide whether it is to be able to penetrate through the minute micropyle, or through the firm egg-envelope, into the egg, there to effect fertilization. An individual with less well formed sperm-cells will be able to fertilize fewer eggs, and therefore to leavefewer descendants which might inherit its tendency to produce inferior sperm-cells, and conversely. Thus it is not the sperm-cells of any one individual which are selected according to their fitness, it is the individuals themselves which compete with one another in the production of germ-cells which shall fertilize best, that is, most certainly. The struggle is thus not intercellular, but a struggle between persons.

The same is true of all cells differentiated for particular functions; every new kind of glandular, muscular, or nerve cell, such as have arisen a thousandfold in the course of phylogeny, can only have resulted from a struggle between individuals which turned on the possession of the best cells of a particular kind,not from a struggle between the cells themselves, since these would gain no advantage from serving the organism, as a whole, better than others of their kind. In regard to the sex-cells we might admit, in addition to personal selection, the possibility of an internal struggle between the sperm-cells or egg-cells of the same individual, inasmuch as each of these cells is the primordium of a new individual, and as those better adapted for reproduction might transmit their better quality to these new individuals. I will not here enter into my reasons for regarding this idea as erroneous, for in any case this interpretation would not apply to any other kind of cells. If, for instance, it were a question of the transformation of an ordinary mucus or salivary gland into a poison gland, it would not matter in the least to the individual cell whether it yielded a harmless or a poisonous secretion; but individuals with many poisonous cells would have an advantage in the struggle for existence.

I agree so far with Plate when he refers the differentiation of the tissues entirely to personal selection, but not in his further conclusion that histonal selection does not exist. The ground-plan of the architectural structure of the organ depends upon personal selection, but the realization of the plan in particular cases is not predetermined down to the minutest details, but is regulated by histonal selection, and is thus to a certain extent an adaptation to local conditions of stimulus. The direction, strength, and size of every single bone lamella is not predetermined from the germ, but only the occurrence and nature of bone-cells and bone lamellæ in general. The direction and the strength which these bone lamellæ may assume depends on the local conditions of strain and pressure which affect the cell-mass, as is shown very clearly by the spongiosa of an obliquely healed bone, which we have already described.

But let us now turn to the question which is here most important for us:whether functional adaptations can be transmitted. We mustadmit that the insight we have so far gained into the causes of these adaptations does not make it much easier to answer the question. Histonal selection is a purelylocalprocess of adaptation to the conditions of stimuli prevailing at the moment, and no one will be likely to suppose that the distorted position of the spongiosa of a badly healed fracture could reappear in the straight bone of a descendant; this would be quite contrary to the principle, for the crooked lamellæ would in that case no longer be the best adapted. Even the questionwhether the strengthening of a muscle through use can be transmittedcannot be answered in the light of the knowledge we have hitherto gained. The 'trophic effect of the functional stimulus' is brought into activity through entirely local influences, through which only the parts most strongly affected by the stimulus can be caused to vary. Thus the problem remains unaltered, How can purely local changes, not based in the germ, but called forth by the chances of life, be transmitted to descendants?

If all species, even in the highest groups, reproduced by dividing into two, we might imagine that a direct transmission of the changes acquired in the course of the individual life through use or disuse took place, though this would presuppose a much more complicated mechanism than is apparent at first sight. But it is well known that multiplication by fission is for the most part restricted to simple organisms, and that the great majority of modern plants and animals reproduce by means of germ-cells, which develop within the organism in regions often very remote from the parts, the results of the exercise of which are said to be transmitted. Moreover, the germ-cells are of very simple structure, at least as far as our eyes can discern; for we see in a germ-cell neither muscles nor bones nor ligaments, glands nor nerves, but only a cell-body consisting of that semifluid living matter to which the general name of protoplasm has been given, and of a nucleus, in regard to which we cannot say that it differs in any essential or definite way from the nucleus of any other cell. How then could the changes which take place in a muscle through exercise, or in the degeneration of a joint in consequence of disuse, communicate themselves to a germ-cell lying inside the body, and do so in such a fashion that this germ-cell is able, when it grows into a new organism, to produce of itself, in the relevant muscle and joint, a change the same as that which had arisen in the parent through use and disuse? That is the question which forced itself upon me very early, and in following it up I have been led to an absolute denial of the transmission of this kind of 'acquired characters.'

In order to explain how I reached this result, and what it isbased upon, it is indispensable that we should first make ourselves acquainted with the phenomena of heredity in general, and the inseparably associated phenomena of reproduction, so that we may form some sort of theoretic conception of the process of inheritance—a picture, necessarily provisional and imperfect, of the mechanism which enables the germ-cell to reproduce the whole organism, and not merely, like other cells, others like itself. We are thus led to an investigation of reproduction and heredity, at the conclusion of which we shall feel justified in returning to the question of the inheritance of acquired characters, in order to give a verdict as to the retention or dismissal of the Lamarckian principle.

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